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Giant Water Scavenger Beetles Hydrophilus Subgenus Dibolocelus (Coleoptera: Hydrophilidae) from Mexico with Description of Two New Species

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Giant Water Scavenger Beetles Hydrophilus Subgenus Dibolocelus (Coleoptera: Hydrophilidae) from Mexico with Description of Two New Species Zootaxa 5027 (3): 387–407 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2021 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.5027.3.5 http://zoobank.org/urn:lsid:zoobank.org:pub:0B8E6B3B-3F14-4959-AB39-ADF60E98EE43 Giant water scavenger beetles Hydrophilus subgenus Dibolocelus (Coleoptera: Hydrophilidae) from Mexico with description of two new species ROBERTO ARCE-PÉREZ1, EMMANUEL ARRIAGA-VARELA2*, RODOLFO NOVELO-GUTIÉRREZ1 & JOSÉ L. NAVARRETE-HEREDIA2 1Instituto de Ecología, A.C., Red de Biodiversidad y Sistemática, Carretera Antigua a Coatepec, 351, El Haya, 91070, Xalapa, Veracruz, Mexico. [email protected]; https://orcid.org/0000-0002-7714-6943 [email protected]; https://orcid.org/0000-0003-3163-2820 2Centro de Estudios en Zoología, CUCBA, Universidad de Guadalajara, Apdo. postal 134, Zapopan, Jalisco, Mexico. [email protected]; https://orcid.org/0000-0002-3336-8528 *Corresponding author. [email protected]; https://orcid.org/0000-0001-6206-9212 Abstract The Hydrophilus (Dibolocelus) Bedel species from Mexico and adjacent areas are treated on the basis of the study of 142 adult specimens and published data. Two new species are herein recognized and described: H. (D.) nucleoensis Arce-Pérez & Arriaga-Varela sp. nov. from Mexico, Guatemala and Nicaragua, H. (D.) pseudovatus Arce-Pérez & Ar- riaga-Varela sp. nov. from Mexico. Three already known species are re-described and their geographical distribution is updated, H. (D.) ovatus Gemminger & Harold and H. (D.) pollens Sharp and H. (D.) violaceonitens Jacquelin du Val, a species resurrected as distinct to H. (D.) smaragdinus. Hydrophylus (D.) cf. purpurascens (Régimbart) is also redescribed but their identity is questionable. The lectotype of H. (D.) pollens Sharp is designated. Within Mexico, the state with the highest known diversity is Veracruz with five species. A key to the species of Hydrophilus (Dibolocelus) from Mexico and Central America is provided. Key words: Aquatic beetles, Neotropical region, new taxa, status restored, morphology, distribution Introduction The genus Hydrophilus Geoffroy, 1762 is a group of distinctly large aquatic beetles, with saprophagous or facultative phytophagous habits (Martin Fikáček, personal communication). It is currently composed of 50 species worldwide, classified in three subgenera: Hydrophilus (sensu stricto) Geoffroy, 1762, Dibolocelus Bedel, 1891 and Temnopterus Solier, 1834 (Hansen 1999; Toussaint et al. 2017). The first two are represented in the New World by 19 species: one is found exclusively in the Nearctic, 16 in the Neotropics, and two have their distribution range throughout both zoogeographic regions (Hansen 1999, Short & McIntosh 2014, 2015). Hansen’s (1991) placement of Dibolocelus as a subgenus within Hydrophilus rather than a distinct genus was recently tested and supported by Short (2010) through a morphological phylogenetic analysis of the tribe Hydrophilini that included the study of Hydrophilus species from North and Central America. A molecular phylogeny of tribe Hydrophilini (Toussaint et al. 2017) supported the monophyly of Dibolocelus; however, it seemed to suggest that its inclusion would render Hydrophilus sensu stricto a paraphyletic group. Additional taxonomical sampling is needed in order to have a clear insight of the subgeneric relationships within Hydrophilus. Arce-Pérez & Morón (2013) reviewed the Hydrophilus s.str. species from Mexico and Central America, updating the known distribution of the species, while Short & McIntosh (2014) reviewed the Hydrophilus species of the United States of America and Canada, updating their distribution and providing new records from Mexico. Currently, a total of 10 species of Hydrophilus (Dibolocelus) are known (Hansen 1999; Short & McIntosh 2014, 2015), one of them is distributed in the Nearctic Region and the remaining nine restricted to the Neotropics. Dibolocelus specimens are not abundant in most of the entomological collections of Mexico and Central America. This situation has led to a scarcity of information about their distribution in the region and uncertainty about the Accepted by G. Gustafson: 17 Aug. 2021; published: 1 Sept. 2021 387 presence of certain species. So far, four species are recorded from the area: H. (D.) smaragdinus Brullé, 1837, H. (D.) pollens Sharp, 1887, H. (D.) purpurascens (Régimbart, 1901) and H. (D.) ovatus Gemminger & Harold, 1868 (Hansen 1999; Arce-Pérez & Morón 2013; Short & McIntosh 2014). In this contribution, we update the information about the diversity of this subgenus in Mexico based on the study of specimens deposited in several entomological collections of the country and other Central American countries. The type of H. (D.) pollens deposited in the Mu- seum of Natural History, London, UK, was also examined. We describe two new species and re-describe the known species from the region. An illustrated key for the identification of the known species of the subgenus in Mexico and Central America, and updated distribution maps are provided. Material and methods The present contribution is based on the study of 138 specimens of Hydrophilus (Dibolocelus) deposited in the fol- lowing entomological collections: AOBC A. O. Bachmann personal Collection, Argentina (Axel O. Bachmann); BMNH The Natural History Museum, London, United Kingdom (Christine Taylor, Maxwell Barclay); CNIN Colección Nacional de Insectos, Universidad Nacional Autónoma de México, Mexico City, Mexico (Santiago Zaragoza-Caballero); CZUG Centro de Estudios en Zoología, Universidad de Guadalajara, Zapopan, Mexico (José L. Navarrete-Heredia); INBIO Instituto Nacional para la Biodiversidad, Costa Rica (Ángel Solís); IEXA Colección Entomológica, Instituto de Ecología, A.C. Xalapa, Mexico (Roberto Arce-Pérez); INIFAP Colección Instituto Nacional de Investigaciones Forestales, Agrícolas y Pecuarias, Celaya, Mexico (Antonio Marín Jarillo); MNHN Muséum National d’Histoire Naturelle, Paris, France (Antoine Mantilleri); NMPC National Museum, Prague, Czech Republic (Martin Fikáček); SEMC University of Kansas, Lawrence, Kansas, USA (Andrew E.Z. Short); UVGC Colección Entomológica, Universidad del Valle de Guatemala, Guatemala, Guatemala (Enio Cano). The specimens were studied with a Carl Zeiss IV-B or an Olympus SZX 16 stereomicroscope. The microphoto- graphs were taken with a Nikon DS-U3 camera attached to a Nikon SMZ25 stereomicroscope and a Canon EOS 1100D digital camera attached to an Olympus BX41 compound microscope. Habitus photographs were taken using a Canon EOS 550D digital camera with a Canon MP-E65mm f/2.8 1–5× macro lens attached or a Nikon Coolpix P500. The photographs were later processed in Adobe Photoshop CS5 program. Descriptions are based on male specimens and comments on the sexual dimorphism and interspecific variation are provided. Measurements are in millimetres. Geographical coordinates were obtained in Google Earth (https://www.google.com/earth) and provided in brackets, when they were not part of the original label data in the list of studied material. The maps were gener- ated on SimpleMappr (https://www.simplemappr.net/#tabs=0) and edited with Corel Draw. Results Taxonomy Hydrophilus (Dibolocelus) Bedel, 1891 Dibolocelus Bedel, 1891: 309. Type species: Hydrophilus palpalis Brullé, 1837. Diagnosis. In addition to the characters for the genus (Short 2010), adult Hydrophilus (Dibolocelus) can be differen- 388 · Zootaxa 5027 (3) © 2021 Magnolia Press ARCE-PÉREZ ET AL. tiated from those of Hydrophilus (s. str.) by a more convex and robust body shape; maxillary palps usually modified being generally longer, broadened and curved towards apex, sometimes inflated and curved at mid-length, with third palpomere with deep and wide ventral excavation or flat and slightly widened towards the apex; labial palp widely triangular and inflated, clavate or subparallel in shape, with or without setae on lateral margins; fifth protarsomere wide and longer than any of the previous ones, sometimes slightly flattened and with a ventral laminar expansion, never as large as those in Hydrophilus (s.str.); internal spur of anterior tibia flat, sub-rectangular or rounded, with straight or curved external margins, if sub-rectangular then it can be short or long with its apex straight, irregular or clearly bifid; abdominal ventrites 3 and 4 have slight to well-marked oval, glabrous area at mid-line, and ventrite 5 has a wide oblong or sub-triangular glabrous area; ventrites 3 to 5 slightly raised at midline, flattened or forming a keel, slightly projected above ventrite posterior margin. Hydrophilus (Dibolocelus) ovatus Gemminger & Harold, 1868 Figs. 1, 2; 7–10, 41. Hydrophilus ovalis Ziegler, 1844: 45 (primary homonym of Hydrophilus ovalis Brullé, 1837: 53, and Hydrophilus ovalis Castelnau, 1840: 50). Dibolocelus ovalis (Ziegler); Régimbart, 1901: 225. Hydrophilus ovatus Gemminger & Harold, 1868: 476 (replacement name for Hydrophilus ovalis Ziegler). Stethoxus ovatus (Gemminger & Harold); Bedel, 1891: 314. Hydrous (Dibolocelus) ovatus (Gemminger & Harold); Kuwert, 1893: 83. Dibolocelus ovatus (Gemminger & Harold); Young, 1954: 196. Studied material. United States of America: Louisiana, New Orleans, 26.vii.1959, At light, W. Gibson col. (3 ♂, 2 ♀) [INIFAP]. Diagnosis. This species is distinguished by the following combination of characters: maxillary palpi slender, third palpomere slightly flattened and broadened apically; internal protibial spur sub-rectangular with apex trun-
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